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of the 3 loculi of the 3-grooved ovary is 1 ovule. The fruit is a
schizocarp and divides into 3 1-seeded, drupe-like fruitlets, which do
not (as in the Geraniaceæ) leave any pronounced column between
them. Endosperm is wanting. The cotyledons are thick and
sometimes slightly coalescent. Tubers often occur.

Fig. 437.—Diagram of
Tropæolum: sp, spur.
Tropæolum.—About 40 species; all from America.
Pollination.—The spur is the receptacle for the nectar; the flowers are
protandrous; the anthers open first, and one by one take up a position in front of
the entrance to the spur, resuming their original position when the pollen is shed;
the stigma finally takes their place after the filaments have bent backwards.—
These plants have an acrid taste (hence the name “Nasturtium,” “Indian Cress”),
on which account the flower-buds and young fruits of T. majus are used as capers.
Some species are ornamental plants.
Order 5. Balsaminaceæ. Herbaceous, chiefly annual plants with
juicy, brittle stems, so transparent that the vascular bundles may be
distinctly seen. The leaves are simple, usually scattered,
penninerved and dentate; stipules are wanting, but sometimes large
glands are present in their place at the base of the petioles. The
flowers are strongly zygomorphic; of their five 5-merous whorls the
petal-stamens are suppressed (S5, P5, A5 + 0, G5); the sepals are
coloured, the 2 anterior ones (Fig. 438 3, 5) are very small or entirely
suppressed, the posterior one is very large and elongated into a
spur, and the 2 lateral ones pushed forward; sometimes the weight
of the spur turns the flower completely round, so that the posterior
leaves assume an anterior position; apparently only 3 petals, since
the lateral and the posterior petals become united in pairs, and the
anterior is larger and differently shaped; the 5 stamens have very
short and thick filaments united at the base, and their anthers finally
adhere together and remain in this condition, covering over the
gynœceum; the filaments ultimately rupture at the base, and the
entire anthers are raised on the apex of the gynœceum as it grows
up. The gynœceum has a sessile stigma and a 5-locular ovary. The
fruit is a capsule which, on maturity, opens suddenly when irritated,
dividing into valves from the base upwards, and as the 5 valves roll
up elastically, the seeds are shot out on all sides to considerable
distances; a central column persists (Fig. 439). The embryo is
straight, and without endosperm.

Fig. 438.—Diagram of
Impatiens glanduligera.
 Fig. 439.—Fruit of Impatiens.
Impatiens; in Europe only I. noli-me-tangere. 225 species; especially from Asia.
Several species have two kinds of flowers: small, cleistogamic, but fertile; and
large, coloured flowers, which in I. balsamine (ornamental plant, E. Ind.) are
protandrous and pollinated by hive-and humble-bees, as they suck the honey from
the spur.
Order 6. Limnanthaceæ. The flowers are regular and differ from all the other
orders in the family by having the carpels not in front of the petals, but in front of
the sepals (which are valvate), and further, the loculi are nearly free individually,
but with a common gynobasic style; the ovules are ascending and apotropous
(anatropous with ventral raphe). The fruit is a schizocarp, with nut-like cocci.—
Limnanthes (4 species; N. Am.) perhaps belongs to another family.
Order 7. Humiriaceæ. Trees and shrubs; about 20 species; Trop. Am.

Family 13. Columniferæ.

The chief characteristics of the orders belonging to this family are
the ☿, regular, generally 5-merous, hypogynous flowers with 5-
merous calyx, sepals united and valvate in the bud; petals 5, free
(often twisted in the bud); stamens ∞ e.g.: 10, in two whorls, but one
of these is more or less suppressed, often altogether wanting, or
replaced by 5 staminodes, while the other (inner whorl) is generally
divided more or less deeply into a large number of anther-bearing
filaments. The filaments too (except Tiliaceæ) are united into a tube,
which, especially in the Malvaceæ, forms a long column in the centre
of the flower, surrounding the gynœceum (Figs. 445, 448); in this
case, which is the most pronounced, the filaments are united into
one bundle (monadelphous), in other instances, polyadelphous. The
number of carpels varies greatly (2 to about 50), but they are nearly
always united and form a syncarpous multilocular gynœceum.—The
vegetative characters also closely agree, the leaves are always
scattered and generally stipulate; all the green portions very often
bear stellate hairs, and the bark in all the 3 orders is rich in tough
bast. Mucilage is often present in cells or passages.—This family is
connected with the Ternstrœmiaceæ, from which it is very hard to
draw a sharp line of demarcation, and it is also allied to the Cistaceæ
and to the Gruinales.
Order 1. Sterculiaceæ (including Buettneriaceæ). This is, no
doubt, the least modified order, and one in which the stamens occur
undivided. Obdiplostemonous. The 10 stamens in two whorls are
most frequently united at the base into a short tube, and have 4-
locular, extrorse anthers. The calyx-stamens are nearly always
simple, tooth-like staminodes, situated on the edge of the tube, or
are entirely suppressed. The same relation is found, for instance, in
the Ampelidaceæ and Rhamnaceæ, namely 5 stamens in front of the
5 petals; not infrequently the 5 stamens are doubled (Fig. 441).
Unisexual flowers are found in Sterculia, Cola, Heritiera. The corolla
is often wanting, or developed in an unusual manner. Each loculus of
the ovary (generally 5) always contains more than one ovule. Fruit a
capsule. Androgynophore often present (Helicteres; Sterculia, etc.).
Hermannia, Mahernia, Melochia, etc., have flat petals with twisted æstivation; 5
undivided stamens, which usually are but slightly united at the base, and most
frequently, without staminodes. Thomasia; Helicteres; Sterculia (free follicles).—
Theobroma, Rulingia, Buettneria, Commersonia, Guazuma, etc., have petals
concave at the base, and terminating in a limb abruptly bent back, and at the
boundary between them most frequently ligular outgrowths, as in certain genera of
the Caryophyllaceæ; stamens 5–15–∞, anthers at the edge of a short tube and 5
linear staminodes (Fig. 441).—The Cocoa-tree (Theobroma), (Fig. 440) bears
large, reddish-yellow, berry-like fruits, resembling short cucumbers, but ultimately
becoming leathery to woody; in each of the 5 loculi are 2 (apparently only 1) rows
of horizontal, oily seeds, as large as almonds. Cotyledons large, thick, and
irregularly folded. Endosperm absent (Fig. 442).
49 genera, with about 750 species; almost entirely confined to the Tropics; none
in Europe or in N. Asia.—The seeds of the Cocoa-tree (T. cacao, bicolor, glaucum,
etc., natives of Trop. Am., especially north of the Equator) are used for chocolate
and are also officinal (“Cocoa-beans,” “Cocoa-butter,” “Oil of Theobroma”).
Theobromine. Cola acuminata, Africa.

Fig. 440.—Theobroma cacao. Branch with flowers and fruits (⅙).

 Figs. 441–442.—Theobroma
cacao.

Fig. 441.—Diagram of the flower:

st barren stamens.

Fig. 442.—B Seed in transverse section: n hilum. A

Embryo after the removal of one of the cotyledons.
 Order 2. Tiliaceæ. This differs from the other orders of the
Columniferæ chiefly in the stamens being entirely free from each
other, and also divided into many filaments, as far as the base, or at
all events very far down, so that the flower appears to have
numerous stamens or to be slightly polyadelphous (Fig. 443); in
addition to this, it may be observed that the anthers are 4-locular and
introrse. In Luehea the groups of stamens alternate with the petals.
In a few genera (Corchorus, Triumfetta) 10 free and single stamens
are found in 2 whorls; but, in the majority, groups of free stamens in
separate bundles. The stamens are more or less united in Apeiba,
Luehea. Style simple. Ovary 2-locular. The ovules are pendulous;
raphe turned inwards. The calyx readily falls off; the æstivation of the
entirely free petals is slightly imbricate (not twisted).
 Fig. 443.—Inflorescence of Tilia, with
its winged bracteole (h); a, a axis of the
shoot; the vegetative bud is seen
between the inflorescence and the axis
of the shoot; b petiole of foliage-leaf.
Tilia (Figs. 443, 444). Calyx and corolla 5-merous; the 5 staminal
leaves (opposite the petals) divided as far as the base into a large
number of stamens which are free or united into groups; gynœceum
with 5 loculi in the ovary (opposite the sepals); there are 2 ovules in
each loculus, though the ovary ripens into a 1-seeded nut, which is
not detached from the axis of the inflorescence, but is carried away
by the wind, whirling round and round, its large-winged bracteole
serving as a parachute (Fig. 443).—Only trees, with alternate, obliquely
heart-shaped and dentate leaves; stellate hairs, as in the other Columniferæ, are
often present. The terminal bud of the branch always fails to develop, and the
growth is then continued sympodially by the uppermost axillary buds. The
inflorescence (Figs. 443, 444) is a 3–7-flowered dichasium (Fig. 444 t, d, e),
which is developed in the axil of a foliage-leaf (Fig. 444). The first of its 2
bracteoles (a) is large, thin, leaf-like, and united with the inflorescence, the lower
portion of which forms a broad wing, its so-called “bract”; the second bracteole (b),
on the other hand, remains scale-like, and supports a winter foliage-bud covered
with bud-scales which thus is situated at the base of the inflorescence, and is a
bud of the 2nd order, in relation to the vegetative shoot. This bud is always found
beneath the inflorescence on the branch placed horizontally, and the winged
bracteole is always found above it, a relation which is connected with the fact that
the 2 rows of shoots on the sides of a branch are antidromous with regard to each
other.—The dichasium itself (Fig. 444) terminates with the flower (t); it has 3 floral-
leaves (c, d, e), which soon fall off; c is barren: the other two bear flowers, or few-
flowered dichasia, or unipared scorpioid cymes (indicated in the figure).—The
foliage-leaves are folded in the bud upon the median line (1, 2, 3 in Fig. 444 are
foliage-leaves with their 2 stipules), the inner half is broader than the outer, and
after unfolding is turned away from the mother-axis (the position of the new
inflorescences and vegetative buds is indicated in their axils on the figure).—The
cotyledons on germination appear above the ground as large, lobed leaves.
Of the other genera some have a bell-shaped, gamosepalous calyx, some have
no corolla, the anthers of some open at the apex (Aristotelia, Elæocarpus, etc.),
the majority have a capsule, some have berries, or drupes, some separate into
fruitlets, etc.—Corchorus, Triumfetta (nut, with hooked bristles), Luehea, Apeiba,
etc. Sparmannia is an African genus; 4-merous flowers; fruit a warted capsule;
filaments numerous and sensitive to touch, the external ones are without anthers
and moniliform above. The plant is covered with numerous soft and stellate hairs,
and at the apex of the branches bears several cymose umbels.
 Fig. 444.—Diagram of the inflorescence of Tilia and the
vegetative bud; the position of the leaves is indicated, and
also the position of the inflorescences, which develop from
their axils in the following year.
Pollination in Tilia is effected by insects, especially bees and Diptera, which
swarm round the tree tops, allured by the numerous strongly-scented flowers and
the easily accessible honey (formed in the hollow sepals). As the flowers are
pendulous, the nectar is protected from ruin; and, in addition, the inflorescence is
more or less concealed beneath the foliage-leaf. Self-pollination is impossible, on
account of protandry.—About 470 species (nearly all trees and shrubs); especially
in the Tropics, only a few being found in the temperate, none in the polar regions,
or in high mountainous districts.—The inflorescence of the native species of Tilia is
medicinal. The wood is used for charcoal.—The majority are used for timber, and
for the sake of the bast (“Bast,” “Jute,” the bast of Corchorus textilis, Luehea, and
others).

Order 3. Malvaceæ (Mallows). The plants are easily recognised

by the scattered, simple, palminerved, most frequently lobed,
stipulate leaves, folded in the bud; the perfect, regular, hypogynous
flowers, with gamosepalous, persistent, 5-merous calyx with valvate
æstivation; the 5 petals twisted in the bud and united with one
another at the base, and by the 5 apparently numerous stamens
(Figs. 445, 448), with the filaments united into a tube, with reniform
bilocular anthers opening by a crescentic slit (in 2 valves). Carpels
3–∞ united into one gynœceum; the embryo is curved and the
cotyledons are folded (Figs. 447, 451); endosperm scanty, often
mucilaginous.—Most of the plants belonging to this order are herbs,
often closely studded with stellate hairs. The leaves are most
frequently palmatifid or palmatisect.

Fig. 445.—Longitudinal section through the flower of

Malva silvestris.
 Fig. 446.—Diagram of Althæa rosea: i the
epicalyx.
An epicalyx is often found formed by floral-leaves placed close beneath the
calyx, in some 3, in others several. The median sepal is posterior in the species
without epicalyx, often anterior in those which have an epicalyx.—The petals are
twisted either to the right or to the left in accordance with the spiral of the calyx;
they are most frequently oblique, as in the other plants with twisted corollas, so
that the portion covered in the æstivation is the most developed. The corolla drops
off as a whole, united with the staminal tube.—Only the 5 petal-stamens are
developed, but they are divided into a number of stamens, placed in 2 rows, and
provided only with half-anthers (leaf-segments, see Fig. 446; the sepal-stamens
are completely suppressed); these 5 staminal leaves are then united into a tube,
frequently 5-dentate at the top, and bearing the anthers on its external side. The
pollen-grains are specially large, spherical and spiny. There are from 3 to about 50
carpels united into one gynœceum and placed round the summit of the axis which
most frequently projects between them. There is only 1 style, which is generally
divided into as many stigma-bearing branches as there are carpels (Figs. 445,
448). The fruit is a schizocarp or capsule. Endosperm (Figs. 447 A, 451) scanty,
often mucilaginous round the embryo, which is rich in oil.
The order is the most advanced type of Columniferæ; it stands especially near
to the Sterculiaceæ, but is separated from these and from the Tiliaceæ, among
other characters, by its 2-locular (ultimately 1-chambered) anthers.
 The sub-orders may be arranged as follows:—
I. Carpels in one whorl.
A. The fruit a capsule, most frequently with loculicidal dehiscence, and
many seeds in each loculus.

1. Gossypieæ. The staminal-column is naked at the apex,

blunted, or 5-dentate.—Gossypium (the Cotton plant) has an
epicalyx of 3 large ovate-cordate leaves, an almost entire, low and
compressed calyx. Solitary flowers. Large, most frequently yellow,
corollas. A 3–5-valved capsule with many spherical seeds. “Cotton”
is the seed-hairs developed upon the entire surface of the seeds
(Fig. 447), and consists of long, 1-cellular hairs, filled with air (and
therefore white); these are thin-walled, with a large lumen, and
during drying twist spirally, and come together more or less in the
form of bands. They consist of cellulose, and have a cuticle.—
Hibiscus has several, most frequently narrow, epicalyx-leaves, a
distinct 5-toothed or 5-partite calyx.—Abutilon; Modiola.

Fig. 447.—A Seed of Gossypium with hairs; B the same in

longitudinal section.
2. Bombaceæ. The staminal tube is more or less deeply cleft into bundles,
sometimes almost to the base; pollen smooth, style simple with capitate, lobed
stigma. Almost all plants belonging to this group are trees, and in many instances
have large barrel-shaped stems, that is, swollen in the centre, and sometimes
covered with large warts. The wood is exceptionally light and soft. The flowers are
often enormously large, and have beautiful petals; in some they unfold before the
leaves. The capsule-wall is sometimes closely covered on its inner service with
long, silky, woolly hairs, while the seeds themselves are generally without hairs.
These hairs, however, on account of their brittle nature, cannot be used like those
of the Cotton-plant. Digitate leaves are found in the Baobab-tree (Adansonia) from
Africa, noted for its enormously thick, but short stem, and in the American Silk-
cotton trees (Bombax, Eriodendron, Chorisia). Ochroma, Cheirostemon, Durio,
and others also belong to this group. Durio is noted for its delicious fruits, which
have a most unpleasant smell.
[Bombax malabaricum is diplostemonous; the five sepal-stamens repeatedly
branch, and the filaments bear unilocular anthers; the five petal-stamens bear
bilocular anthers.]

B. Schizocarps, with 1-seeded fruitlets, most frequently nut-like

and reniform (Figs. 449, 451).
3. Malveæ, Mallow Group. The carpels are arranged in one
whorl (Fig. 449); the number of stylar-branches equals that of the
carpels; fruitlets 1-seeded, reniform, indehiscent, but detaching
themselves from one another and from the persistent central column
(Figs. 450, 451).—Malva has an epicalyx of 3 free leaves. A flower with 2
suppressed bracteoles is situated in the axil of the foliage-leaves; one of these
supports a homodromous foliage-shoot which forms a repetition of the main axis,
the other an antidromous flower which continues the branching as a unipared
scorpioid cyme.—Althæa, Rose Mallow, has an epicalyx of 6–9 leaves united at
the base.—Lavatera, Sida, Anoda, Bastardia, etc., have no epicalyx.
 Figs. 448–451.—Malva
silvestris.

Fig. 448.—The flower after

removal of the perianth
(5/1).]

Fig. 449.—The fruit (5/1).

 Fig. 450.—A fruitlet (5/1).

Fig. 451.—The same in

longitudinal section.
4. Ureneæ, have always only 5 carpels arranged in 1 whorl, with 1 ovule in
each loculus, and the fruit a schizocarp, generally with nut-like fruitlets provided
with warts and hooks; but in some they dehisce by 2 valves (capsule). They differ
principally from the other groups in having twice as many stylar-branches as
carpels; the staminal tube is naked at the point, blunt or 5 toothed.—The genera
Urena, Pavonia, Malachra, Malvaviscus (with berry-like fruits) belong to this group.
II. Carpels arranged in a spherical head in five groups opposite to
the petals.
 5. Malopeæ, differ from all the others in having a large number of fruitlets
arranged irregularly in a round head, and separating considerably from each other
even before maturity; there is, however, only 1 style, divided into a corresponding
number of branches (this condition may be considered to have arisen from the
branching [dédoublement] of 5 carpels). Malope has 3 large, heart-shaped
(Kitaibelia 6–9) epicalyx-leaves, united at the base. Palava has no epicalyx.
Pollination. The majority have protandrous flowers, and are pollinated by
insects. Between the basal portions of the 5 petals, there are 5 nectaries,
protected from the rain by hairs, e.g. in Malva silvestris. When the flower first
opens the numerous anthers occupy the centre of the flower, and the still
undeveloped stigmas are concealed in the staminal tube; in the next stage the
anthers are withered and empty, and the stigmas protrude and assume their
places (Fig. 452). The large-flowered forms, it appears, are pollinated only by
insects; but self-pollination takes place in small-flowered forms, as, for example, in
Malva rotundifolia, in which the stylar-branches, twisting themselves, place the
stigmas in between the undeveloped anthers.
 Fig. 452.—Anoda hastata: a the bud just opened, the stigmas
are concealed by the anthers; b fully opened flower in ♂-stage; the
upper stamens are developed first, and then the others in
descending order; the stylar-branches are now visible, and lie bent
back on the staminal column; c all the stamens project upwards,
and all the anthers are open, but the stylar-branches are still bent
back; d the anthers are emptied and the filaments shrunk together,
but the styles have now straightened themselves upwards, and the
stigmas are in the receptive condition.
Distribution. 800 species (63 genera), most of which are natives of the
Tropics, especially America. Althæa and some of the species of Malva are natives
of the temperate regions of the Old World, the latter is also found in North
America. Gossypium is tropical, no doubt especially Asiatic (G. herbaceum from
India; G. arboreum from Upper Egypt). Cotton was introduced into Greece in the
time of Herodotus, and was cultivated in America before the arrival of the
Europeans.
Uses. Pungent and poisonous properties are entirely wanting; mucilage, on the
other hand, is found in abundance in all parts of the plant. Medicinal: the root of
Althæa officinalis, leaves and flowers of Malva-species (M. silvestris vulgaris and
borealis) and Gossypium.—The seeds contain a large quantity of fatty oil, which is
in some cases extracted (Cotton-seeds and others). The seed-hairs of the Cotton
plant are the most important product of the order. The cultivated forms of Cotton
belong to several species: G. barbadense, herbaceum, religiosum, arboreum
(Nankin), hirsutum, and others. According to other botanists, there are only 3
species. Bast is obtained from e.g. Hibiscus cannabinus (Gambo-hemp, Africa),
Paritium tiliaceum and Sida retusa. The fruits of certain species of Hibiscus (e.g.
H. esculentus, from Tropical Africa) are used in tropical countries as a vegetable
before they are ripe.—The colouring matter in the flowers of Althæa rosea, var.
nigra, is used for colouring wines, and hence is extensively cultivated in certain
parts of Europe.—Ethereal oils and sweet-scented flowers are rare; but several
species possess a peculiar musk-like odour (Malva moschata, Hibiscus
abelmoschus, and others).—Many are cultivated as ornamental plants on account
of the large flowers, e.g. Hollyhock (A. rosea, etc.), Lavatera trimestris, Malope
grandiflora and trifida, Malva-species, Hibiscus rosa sinensis, syriaca;
Sphæralcea, etc.

Family 14. Tricoccæ.

The very large order Euphorbiaceæ and three smaller ones
belong to this family. They have in common: unisexual, hypogynous,
frequently regular flowers, the perianth most frequently single, rarely
double, or entirely wanting; there is such a great variety in the
structure and parts of the flower that one only can be cited as the
rule: viz. the simple gynœceum composed of 3 carpels forming a 3-
locular ovary, which is frequently more or less deeply grooved
(hence the name, Tricoccæ); in the inner angles of the loculi are
found 1 or 2 (never several) pendulous (except Empetraceæ),
anatropous ovules, with upward and outwardly turned, frequently
swollen, micropyle (Fig. 455). The seed most frequently has a large
endosperm and a straight embryo (Figs. 455 B, 464).—The family
approaches the nearest to the Gruinales and Columniferæ; it may perhaps be
regarded as an offshoot from the Sterculiaceæ.
Order 1. Euphorbiaceæ. Flowers unisexual. In each of the loculi
of the ovary, generally 3, there are 1 or 2 pendulous ovules with
upward and outwardly turned micropyle. The placenta protrudes
above the ovules (Figs. 454, 461 B). On the ripening of the capsule
the 3 carpels separate septicidally, frequently with great violence,
ejecting the seeds and leaving a central column. Endosperm
copious.—For the rest, the flowers present all stages, from genera
with calyx and corolla, to those which are the most reduced in
Nature, namely the naked, 1-stamened flowers of Euphorbia.
The same variety which is found in the flower is also present in
the vegetative parts. Some are herbs, as our Spurges, others are
shrubs and trees; some African Euphorbia-species even resemble
the habit of a Cactus. Leaf-like branches with rudimentary leaves are
found in Phyllanthus (sub-genus Xylophylla) (Fig. 456). The leaves
are scattered or opposite, often stipulate; they are nearly always
simple. Large, highly-branched cells containing a great quantity of
pungent latex are found in many, and watery juice in others. Glands
and glandular hairs are general.—Only a few genera can be
considered in this book.
As an example of the most perfect flowers (which partly reproduce
the Geraniaceous type) may be mentioned, Croton, Manihot, and
Jatropha; 5 sepals, 5 petals, sometimes gamopetalous, andrœcium
diplostemonous, or many-stamened, often monodelphous.

Figs. 453–455.—Ricinus communis.

Fig. 453.—♂-flower (magnified).